This thesis has the overall aims of creating a welfare-sensitive additive index of captive great ape husbandry using the expert opinion of relevant stakeholders, highlighting the aspects of husbandry requiring most improvement and allowing enclosures in different zoos and wildlife parks to be ranked in a uniform way. I investigated the attitudes of experts on the management of captive great apes to gain a greater understanding of the features of their environment that may be critical in maintaining a high standard of welfare. From the experts ranking of the different Levels of facilities offered to the great apes I constructed an additive index with 17 key attributes with between two and four Levels for each. The Great Ape Welfare Index (GAWI) derived from expert opinion has Social structure, Enclosure appearance, Group size, Enclosure furnishings and Avoidance provision as the most important attributes of a management system. The GAWI was then validated by recording the behaviour of the captive great apes housed in Australian and overseas zoos, and comparing their activity budgets to wild individuals. Using a Penrose Distance Statistic a significant negative correlation was found between Index score and the departure from wild activity budgets. Additionally I found a significant negative correlation between the change in the proportion of time spent resting and the welfare Index score achieved indicating that those great apes housed in enclosures achieving higher Index scores are more active in general. Finally the husbandry and management systems for great apes currently used by zoos globally were investigated and their ranking in the Index were calculated. This allowed the particular aspects of great ape management requiring improvement to be identified for each enclosure. Comparisons of regional means showed that Australian zoos achieved significantly higher Index scores than those in other parts of the world. The proportion of great apes housed in New Zealand zoos reported as displaying an abnormal behaviour was significantly higher than all other regions with the proportion of great apes housed in Australia displaying significantly less abnormal behaviour when compared to the US. The highest index achieved by any enclosure was 97.5 for an enclosure that housed a group of gorillas, with the lowest index allocated was 52.2 for an orangutan enclosure. The presence of abnormal behaviours was also investigated with the most common abnormal behaviours reported being regurgitation and re-ingestion (15.8%), hair plucking (10.2%) and begging (7.9%) with some animals reported as exhibiting four different abnormal behaviours. The GAWI has the potential to improve the captive environments provided to great apes through highlighting the most critical aspects of their husbandry requiring improvement. This should encourage zoos to aim higher than just complying with minimum standards set by their regional zoological associations. As the index correlates with the behaviour of the great apes housed, their behaviour may more closely resemble wild individuals if improvements are made. The GAWI has been compared to two alternative models to investigate whether the methods used to create the index could be simplified while still allowing adequate discrimination between enclosures and species. This index can be adapted for use in other institutions housing great apes and can act as a model to create indices for other species. The index has been submitted to the primate Taxon Advisory Group of the Australian Regional Association of Zoological Parks and Aquaria for their consideration of its use in their accreditation scheme or husbandry guidelines.

This thesis has the overall aims of creating a welfare-sensitive additive index of captive great ape husbandry using the expert opinion of relevant stakeholders, highlighting the aspects of husbandry requiring most improvement and allowing enclosures in different zoos and wildlife parks to be ranked in a uniform way. I investigated the attitudes of experts on the management of captive great apes to gain a greater understanding of the features of their environment that may be critical in maintaining a high standard of welfare. From the experts ranking of the different Levels of facilities offered to the great apes I constructed an additive index with 17 key attributes with between two and four Levels for each. The Great Ape Welfare Index (GAWI) derived from expert opinion has Social structure, Enclosure appearance, Group size, Enclosure furnishings and Avoidance provision as the most important attributes of a management system. The GAWI was then validated by recording the behaviour of the captive great apes housed in Australian and overseas zoos, and comparing their activity budgets to wild individuals. Using a Penrose Distance Statistic a significant negative correlation was found between Index score and the departure from wild activity budgets. Additionally I found a significant negative correlation between the change in the proportion of time spent resting and the welfare Index score achieved indicating that those great apes housed in enclosures achieving higher Index scores are more active in general. Finally the husbandry and management systems for great apes currently used by zoos globally were investigated and their ranking in the Index were calculated. This allowed the particular aspects of great ape management requiring improvement to be identified for each enclosure. Comparisons of regional means showed that Australian zoos achieved significantly higher Index scores than those in other parts of the world. The proportion of great apes housed in New Zealand zoos reported as displaying an abnormal behaviour was significantly higher than all other regions with the proportion of great apes housed in Australia displaying significantly less abnormal behaviour when compared to the US. The highest index achieved by any enclosure was 97.5 for an enclosure that housed a group of gorillas, with the lowest index allocated was 52.2 for an orangutan enclosure. The presence of abnormal behaviours was also investigated with the most common abnormal behaviours reported being regurgitation and re-ingestion (15.8%), hair plucking (10.2%) and begging (7.9%) with some animals reported as exhibiting four different abnormal behaviours. The GAWI has the potential to improve the captive environments provided to great apes through highlighting the most critical aspects of their husbandry requiring improvement. This should encourage zoos to aim higher than just complying with minimum standards set by their regional zoological associations. As the index correlates with the behaviour of the great apes housed, their behaviour may more closely resemble wild individuals if improvements are made. The GAWI has been compared to two alternative models to investigate whether the methods used to create the index could be simplified while still allowing adequate discrimination between enclosures and species. This index can be adapted for use in other institutions housing great apes and can act as a model to create indices for other species. The index has been submitted to the primate Taxon Advisory Group of the Australian Regional Association of Zoological Parks and Aquaria for their consideration of its use in their accreditation scheme or husbandry guidelines.

Although human biomedical and physiological information is readily available, such information for great apes is limited. We analyzed clinical chemical biomarkers in serum samples from 277 wild- and captive-born great apes and from 312 healthy human volunteers as well as from 20 rhesus macaques. For each individual, we determined a maximum of 33 markers of heart, liver, kidney, thyroid and pancreas function, hemoglobin and lipid metabolism and one marker of inflammation. We identified biomarkers that show differences between humans and the great apes in their average level or activity. Using the rhesus macaques as an outgroup, we identified human-specific differences in the levels of bilirubin, cholinesterase and lactate dehydrogenase, and bonobo-specific differences in the level of apolipoprotein A-I. For the remaining twenty-nine biomarkers there was no evidence for lineage-specific differences. In fact, we find that many biomarkers show differences between individuals of the same species in different environments. Of the four lineagespecific biomarkers, only bilirubin showed no differences between wild- and captive-born great apes. We show that the major factor explaining the human-specific difference in bilirubin levels may be genetic. There are human-specific changes in the sequence of the promoter and the protein-coding sequence of uridine diphosphoglucuronosyltransferase 1 (UGT1A1), the enzyme that transforms bilirubin and toxic plant compounds into water-soluble, excretable metabolites. Experimental evidence that UGT1A1 is down-regulated in the human liver suggests that changes in the promoter may be responsible for the human-specific increase in bilirubin. We speculate that since cooking reduces toxic plant compounds, consumption of cooked foods, which is specific to humans, may have resulted in relaxed constraint on UGT1A1 which has in turn led to higher serum levels of bilirubin in humans.

No description available.

Unsustainable international wildlife trade together with destruction and pollution of wild nature and spread of the invasive species, has long been recognised as a major threat to biodiversity. The subject of this thesis is an evaluation of international trade with some groups of mammals (Great apes and Rhinos) covered by CITES convention between European Union and third countries according EU Annual Reports in 2000 ? 2009. The trade has been divided into export and import and within these categories were evaluated main traded species, types of specimens, exporter and importer, source and purpose of the trade. It was found the trends of Great apes export and import increased, trend of Rhinos export decreased, the trend of Rhinos import increased over this period.

Increasing anthropogenic alterations propelled by a growing human population paired with ecological perturbations and climate change has amplified rates of disease transmission at the human-wildlife interface. While attention has focused primarily on diseases that cause high rates of morbidity and mortality, there is a dearth of research on more common, non-lethal “mild” infections. However, despite less obvious and immediate consequences, these infections still have long-term effects on both public health and the conservation of wildlife. Currently, disease research is primarily cross-sectional, with a lack of longitudinal studies, leading to an undervaluation of the dynamic nature of disease systems. In addition to pathogen monitoring, concurrently being able to measure immune system activation will help to clarify the effects of non-lethal diseases on host health and to provide further insights into life-history trade-offs. Here, I investigated malaria parasite (Plasmodium spp.) infections, a “mild” disease, in wild habituated chimpanzees (Pan troglodytes verus) residing in Taï National Park (TNP), Côte d'Ivoire. I used historical biological samples collected from non-human primates (including chimpanzees) and humans, as well as collected mosquitoes within their habitat. First, I identified longitudinal patterns of malaria parasite prevalence detected in chimpanzee faeces; next, I validated a biomarker of immune system activation, urinary neopterin, in wild chimpanzees; and lastly, within a larger ecological framework, I examined the interface of malaria parasite transmission between humans and non-human primates sharing a habitat. With a longitudinal study design, I found substantial intra- and inter-annual fluctuations in the faecal detection of malaria parasites across four non-consecutive sampling periods between 2004 and 2015. Peaks were observed during wet seasons—suggesting that environmental factors relating to vector abundance determine infection patterns. A higher prevalence was also detected in younger individuals, suggesting that the availability of susceptible hosts plays a role. With variations in parasite detection, similar trends should also be observed in health status. Urinary neopterin, an early inflammation marker of the non-specific immune response, increases during malaria parasite infections in humans and has been validated as a marker of immune system activation in laboratory and captive non-human primates. However, it was unclear whether it would be sensitive enough to provide a clear signal in mild diseases against the back-drop of co-infections commonly seen in wildlife. Therefore, we first needed to validate urinary neopterin as a biomarker of immune system activation during severe disease in wild animals. I measured urinary neopterin before, during, and after a severe respiratory outbreak and showed that levels corresponded to respiratory symptoms and predicted mortality. While urinary neopterin is sensitive enough to detect changes in immune system activation during severe disease, future research should still aim to validate its use in mild diseases, such as malaria. Finally, human-to-animal disease transmission is known to occur in TNP, with direct declines in chimpanzee populations observed that resulted from several outbreaks caused by human respiratory diseases. Given the zoonotic origin of malaria parasites in humans, I examined the genetic diversity of malaria parasites infecting humans and non-human primates sharing a habitat. Mosquitoes were also captured to identify potential vectors that may bridge transmission between host species. Only P. malariae was found in both humans and chimpanzees—however, the directionality of cross-species transmission would require a larger sample size to correctly assess. Additionally, no anopheline mosquitoes, the only known vector of mammalian malaria parasites, or mosquitoes positive for human- or great ape-specific malaria parasites were captured—suggesting that transmission events may be rare due to the sparsity of vectors in this region. This thesis shows that malaria epidemiology is a temporally and spatially diverse system that requires the use of longitudinal datasets and diverse sampling schemes. This thesis provides a baseline of data on which future malaria parasite research can build. Additionally, the validation of urinary neopterin will allow researchers to pursue questions on how mild diseases affect host health and to investigate questions relating to strategies and variations in life history trade-offs. This thesis is relevant for research on wildlife disease ecology, eco-immunology, and in the creation of pathogen and health surveillance programs.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第20962号 / 理博第4414号 / 新制||理||1634(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)教授 湯本 貴和, 教授 古市 剛史, 教授 友永 雅己 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

Orangutans serve as an important species in hominoid studies given their relatedness to the genus Homo, a flagship taxon chosen to represent forest conservation efforts in Indonesia, and an intriguing model for studying alternative reproductive tactics. Despite interest in the species and their importance to their environment, orangutans are exceptionally challenging to study in the wild and captivity. In this dissertation, I investigated the life history of great apes under human care and the growth/ developmental patterns of orangutans. Studbooks, records of all births and deaths of captive animals, compose the largest life history datasets for great apes. I compared the following life history variables for the four great ape species housed in the United States: age at first successful reproduction, interbirth intervals, prenatal death rate, twinning rate, number of offspring, age at last reproduction, and lifespan. Results showed species-specific differences in median age at first birth among females, age at first successful fertilization among males, and number of offspring sired by males and produced by females. The four great ape species had similar ages at last birth among females, age at last successful fertilization among males, and interbirth intervals. Overall, I found that great apes have similar reproductive stages of their lives and capabilities when living with low ecological risk despite some of the differences that exist in their life history parameters. These findings can be used in comparative studies of humans and their most recent living biological ancestors. Next, I investigated the process of flanging in male orangutans. Male bimaturism is one of the most intriguing aspects of orangutans’ biology, yet there is still much that is unknown about the flanging process. All males begin as an unflanged male with the potential to flange at a random timepoint in their lifetime. I conducted a longitudinal study of two males, Budi who flanged and Kembali who remained unflanged, housed at the Toronto Zoo. For Budi, flanging lasted approximately two years. In this time, he increased in body size by approximately 78%, increased his testosterone levels, grew cheek flanges, and began long calling. Throughout flanging, his cortisol levels also increased and his C-peptide of insulin levels fluctuated, both signs of the energetic cost associated with flanging. These findings highlight the extent to which flanging affects male orangutans in terms of their morphology, physiology, and behavior. Lastly, I investigated differences in muscle mass amongst the three adult morphs of orangutans (females, unflanged males, flanged males). Adult unflanged males are approximately the same body size as adult females whereas adult flanged males are approximately twice the size of adult unflanged males and females. Thus, I predicted flanged males would have the greatest estimated lean body mass (ELBM) followed by unflanged males and females. To test this hypothesis, I analyzed creatinine and specific gravity in a large dataset of urine samples from orangutans under human care from zoos in the United States and a smaller dataset from wild Bornean orangutans at Gunung Palung National Park (GPNP) in West Kalimantan, Indonesia. While my predictions were supported by mean ELBM values, I did not find that the different age-sex classes of adult orangutans were statistically different from one another. When we analyzed both datasets together, however, we found that flanged males had significantly higher ELBM compared to all other age-sex classes and that wild orangutans have significantly lower ELBM compared to their captive counterparts. Given the variation in body size amongst age-sex classes, I believe that sex differences in muscle mass will be distinguishable with a larger sample size. Collectively, these findings in my dissertation highlight differences amongst the life history patterns of great apes, which have classically been viewed as being largely similar to one another, and the distinctiveness of having three adult morphs in a primate species. / 2025-06-27T00:00:00Z

abstract: Early weaning, slow somatic and dental growth, and late age at reproduction are all part of a suite of energetic trade-offs that have shaped human evolution. A similar suite of energetic trade-offs has shaped the evolution of the indriid-palaeopropithecid clade, though members of this clade exhibit extremely fast dental development and nearly vestigial deciduous teeth. The development and functional occlusion of the primary postcanine dentition (i.e., deciduous premolars and molars) coincides with several life history parameters in great apes and indriids. This dissertation explored great ape dental macrowear, molar development in indriids, and molar size in lemurs with a broader goal of improving reconstructions of life history profiles in extinct primates. To this aim, macrowear and dental development were analyzed in apes and lemurs, respectively. Occlusal casts (six great ape species; N=278) were scanned to track mandibular fourth deciduous premolar (dp4) macrowear. Utilizing dental topographic analyses, changes in occlusal gradient and terrain were quantified. A subset of the great ape data (four species; n=199) was analyzed to test if differences in dp4 wear correlate with age at weaning. Using dental histology, molar development was reconstructed for Indri indri (n=1) and Avahi laniger (n=1). Life history and molar size data were collected from the literature. The results of this dissertation demonstrate that most great apes exhibited evidence of topographic maintenance, suggesting dp4s wear in a manner that maintain functional efficiency during growth and development; however, the manner in which maintenance is achieved (e.g., preservation of relief or complexity) is species specific. Dp4 macrowear is not correlated with age at weaning in great apes and is probably unreliable to reconstruct age at weaning in hominins. The pace of molar development in members of the indriid- palaeopropithecid clade did not correlate with body or brain size, an association present in several other primates. Associations of molar size with age at weaning suggest that expanding other developmental models (e.g., the inhibitory cascade) to life history is worth consideration. The broad variation in macrowear, dental development, and size highlights how the primary dentition may correlate with different life history parameters depending on the species and ecological setting, an important consideration when using teeth to reconstruct life history profiles. / Dissertation/Thesis / Doctoral Dissertation Anthropology 2016

Le degré de rétention de l’arboricolisme dans le répertoire locomoteur des hominines fossiles du Pliocène est toujours matière à débat, les études ayant principalement porté sur la courbure des phalanges et la proportion des membres. Vu la récente découverte de DIK-1-1 (A. afarensis) et de la scapula qui lui est associée, l’étude de cet os d’un point de vue fonctionnel est intéressante, puisqu’il est directement impliqué dans la locomotion de presque tous les hominoïdes. Le but de cette étude est de tenter d’établir un lien entre l’orientation supéro-inférieure (SI) et antéro-postérieure (AP) de la cavité glénoïde de la scapula et les comportements locomoteurs chez les grands singes et l’humain moderne. Des analyses comparatives sur les adultes ont été réalisées pour 1) voir s’il existe des différences dans la morphologie étudiée entre les espèces et 2) voir si ces différences peuvent être expliquées par la taille corporelle. Des analyses ontogéniques ont aussi été réalisées pour voir si un accroissement de la taille corporelle pendant le développement et les changements locomoteurs qui y sont associés correspondent à un changement d’orientation de la cavité glénoïde. Les résultats montrent que les humains ont une cavité glénoïde qui est orientée moins supérieurement que les grands singes, mais que Pongo, bien qu’étant le plus arboricole, n’a pas l’orientation la plus supérieure. Les « knuckle-walkers » (Pan et Gorilla) se distinguent des autres hominoïdes avec une orientation de la surface glénoïde relative à l’épine plus inférieure. La taille corporelle ne semble pas influencer la morphologie étudiée, sauf parfois chez le gorille. Seuls l’humain et les mâles Pongo montrent un changement ontogénique dans l’orientation de la cavité glénoïde relativement à l’épine. Sur la base de ces résultats, l’orientation de la cavité glénoïde semble refléter partiellement la fonction du membre supérieur dans la locomotion, mais des recherches plus poussées sont nécessaires. Mots-Clés : Scapula, cavité glénoïde, grands singes, humains, locomotion, arboricolisme. / The degree to which Pliocene fossil hominins have retained some form of arborealism in their locomotor repertoire is still matter of debate, in part because studies linking upper limb morphology to locomotor behaviors in primates mostly focused on phalangeal curvature and limb proportions. Given the recent discovery of DIK-1-1 (A. afarensis) and its associated scapula, investigation of this bone from a functional perspective seemed of interest since it is directly involved in almost every hominoid’s locomotion. The purpose of this study is to try to establish a link between the superoinferior and anteroposterior orientation of the glenoid cavity of the scapula and locomotor behaviors in living great apes and modern humans. Comparative analyses were performed on adult individuals to 1) establish if there were differences across species and 2) verify that those differences, if any, could be linked to overall body size. Ontogenetic analyses were also performed at the intraspecific level to see if a change in body size during development, which is often associated with changes in locomotor behaviors, is related to a change in the orientation of the glenoid cavity. These results show that humans have a more inferiorly oriented glenoid cavity than great apes, but Pongo, even if it is the most arboreal species, does not have the most superiorly oriented glenoid cavity. Knuckle-walkers (Pan and Gorilla) differ from other hominoids, exhibiting a more inferiorly oriented glenoid cavity relative to the spine. Body size does not seem to influence the orientation of the articulation, with a few exceptions for gorillas. Only humans and male Pongo show a significant ontogenetic change in the orientation of the glenoid cavity relative to the spine. On the basis of these results, the orientation of the glenoid cavity seems to reflect only in part the use of the upper limb in locomotion, however, it will need to be investigated further.